Stem gall of Michelia champaca L. (Magnoliaceae) … flower locally known as ‘sampige’ or ‘champa’ has great significance in Hindu mythology and has a number of cosmetic, medicinal

www.tropicalplantresearch.com 90 Received: 22 March 2015 Published online: 30 June 2015

ISSN (E): 2349 – 1183

ISSN (P): 2349 – 9265

2(2): 90–100, 2015

Research article

Stem gall of Michelia champaca L. (Magnoliaceae) induced by

Podothrips sp.: Identification, histochemical and

phytochemical studies

Monnanda Somaiah Nalini1*, K. E. Shilpa

1 and Sekharappa Basavarajappa

2

1Department of Studies in Botany, University of Mysore, Manasagangotri, Mysore–570 006, Karnataka, India 2Department of Studies in Zoology, University of Mysore, Manasagangotri, Mysore–570 006, Karnataka, India

*Corresponding Author: [email protected] [Accepted: 18 June 2015]

Abstract: Galls are organized structures of plant tissues induced by the insects. The present study

deals with the identification of gall-maker on the stem/twigs of Michelia champaca. The gall-

maker was identified as belonging to the order Thysanoptera, and was confirmed as the genus

Podothrips. Histochemical staining was carried out in different portions of gall as well as healthy

twigs for the presence of alkaloids, fats, starch, tannins and proteins. The staining varied in

different tissues. The galls at different development stages (young and mature) were tested for the

presence of total phenolic as well as carbohydrate content. The results revealed high phenolic

content in young galls where as low amount was found in a mature gall tissue, however, a 2.3-fold

difference was noted between the two. Very high concentration of carbohydrate content was

detected in young gall tissues. The gall serves both as a shelter and a food source, to the gall-

maker. The inner walls of the galls are moist and are usually rich in phenolics and carbohydrates.

Our work reports for the first time Podothrips sp. as gall-makers. Further studies on the ecology

of the insect would relate its occurrence on other host species and distribution.

Keywords: Gall - Michelia champaca - Histochemistry - Phenolics - Carbohydrate

[Cite as: Nalini MS, Shilpa KE & Basavarajappa S (2015) Stem gall of Michelia champaca L. (Magnoliaceae)

induced by Podothrips sp.: Identification, histochemical and phytochemical studies. Tropical Plant Research

2(2): 90–100]

INTRODUCTION

Michelia champaca L. is a medium to big sized tree with glossy leaves and yellow or orange flowers of the

family Magnoliaceae, native to the tropical and sub-tropical South and Southeast Asia, including Southern

China. The flower locally known as ‘sampige’ or ‘champa’ has great significance in Hindu mythology and has a

number of cosmetic, medicinal and economic uses. Fresh flowers are extracted into perfumes and medicinal

products which are used as cure for coughs and rheumatism. Cosmetic products such as Joy, J’adore and Dior

contain M. champaca fragrant extracts as ingredients in their composition (Armiyanti et al. 2010).

Michelia champaca is affected by biotic and abiotic factors. Among the biotic factors affecting the plants is

the presence of galls on the plant parts. Galls are abnormal growths on plants and are better understood as a suite

of adaptations to the inducing insect (Raman 2003). They are highly regulated growth manifestations on plants,

modified, symmetrical, natural-plant structures that are the outcome of messages from the inducing insects.

These structures develop as an extension of the host-plant phenotype (Raman 2012). Galls result from the

feeding of living organisms, such as insects, mites, and or mechanical stimulus. The attack of each species

results in a distinctive deformity on leaves, twigs or stem of a host plant. Such response also increases the

plant’s production of growth hormones such as auxins, cytokinins and gibberellins (Caron 2004).

Thrips are one class of well-known gall-forming insects, and belong to the order Thysanoptera. Thrips are

involved in several types of host relationship with plants, and their presence have been documented

(Ananthakrishnan & Raman 1979). Thrips (order-Thysanoptera) are tiny, slender insects with fringed wings.

They are very small, yellow, brown or black, slender insects ranging from 1/16 to 1/8 inch in length. Adults and

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larval thrips feed using a punch and suck technique. Their life cycle includes an egg stage, two larval instars,

two pupal stages, and an adult stage (Buss 2011).

Gall tissues are known to be sources of enzymes, phytochemical compounds such as alkaloids, fats, tannins,

proteins, starch, phenolics, carbohydrates etc. These accumulate in the tissues during gall development. Plant

phenolic compounds play an important role at different levels during plant-microbe interactions. A completely

different action of plant phenolic compounds lies in the defense of plants against pathogen attack (Vereecke et

al. 1997). Histochemistry is the branch of histology dealing with the identification of chemical components of

cells and tissues. Starch deposition occurs widely in the plant body, in the seeds, the parenchyma of the

secondary vascular tissue in the stem. Starch and protein are the principal ergastic substance of the protoplast.

Tannin is the heterogeneous group of phenol derivatives, usually related to glucosides. Fats are widely

distributed in the plant body and they probably occur in small amount of every plant cell. Many woody plants

contain medicinally important secondary products (Marmit & Sharma 2008) which can be analyzed using

staining techniques.

Various types of insect-plant galls exist in nature. Several plant species are reported to possess galls, yet the

occurrence of these in plants is to be documented. So far, no plant galls have been documented from Michelia

species. Since galls on the stem of Michelia was first noted in the Manasagangotri Campus, we undertook the

study to document and identify the gall-inducer and analyse the phytochemical and histochemical differences

between the healthy and galled tissue portions of the stem.

MATERIALS AND METHODS

Collection of the sample and study area

Healthy and galled twigs of Michelia champaca L., were collected from the plants growing in

Manasagangotri campus (12o18' N & 76o12' E), Mysore, during the month of September to December, 2011.

The galled portions were excised from the stem with a metallic plier. The galls were separated according to their

size, as young and mature and photographed.

Identification of the gall-maker from the stem/twigs of Michelia champaca

Various stages of fresh gall materials such as the young and the mature (old) were collected. The collected

tissue was mounted on the clean glass slide and was kept on the field of Steriosome (Leica EZ4, MC234813,

China). The gall tissue was dissected by using clean micro dissection needles and observed for the presence of

the gall-maker. Different stages of insects were photographed by using different magnifications, 8–35X. These

were considered for the documentation studies.

Histochemical comparison between gall and normal tissues

Free hand cut sections of young twigs of Michelia champaca were taken by using clean and sharp blade and

floated in water contained in petriplates. Using pointed brush, fine transparent sections were selected and

immersed and placed in a cleaned glass slide and respective stains were added drop wise and covered with a

clean cover slip. Excess of stain was removed using a tissue paper and observed for the localization of viz.-

starch, tannins, proteins, fats and alkaloids under the microscope at various magnifications (4x, 10x and 40x)

and images were documented and compared.

Preparation of reagents

Reagents were prepared for histochemical studies according to the procedure of Momin & Kadam (2011).

Starch: 0.3 g of iodine and 1.5 g of potassium iodide were dissolved in 100 ml of distilled water. A drop of the

solution was added on the section, left for some time and washed water and observed under microscope.

Protein: Saturated aqueous solution of picric acid is an excellent precipitating agent for protein, staining them

an intense yellow. The sections were allowed to react with the reagent for 24 h. The sections were washed

with 60% alcohol and few drops of aqueous FeCl3 were added.

Tannins: Sections were treated with dilute acidic ferric chloride solution (0.5% to 1% of FeCl3 in 0.1 N HCl),

mounted in clove oil and observed under microscope for the presence of tannins.

Fats: 0.5 g of Sudan III and or Sudan IV is dissolved in 100 ml of 70% alcohol. Sections were immersed in the

stain for 20 min, rinsed quickly with 50% alcohol and mounted in glycerine for observations.

Alkaloids: Wagner’s reagent: One gram of iodine and 2 g of potassium iodide were dissolved in 50 ml of

distilled water.

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Estimation of total phenolic and carbohydrate content in the gall sample

Preparation of the extract: One gram each of young and mature gall samples were finely ground for few

minutes with 10 ml of 70% alcohol by using clean mortar and pestle. The extracts were taken in small

Eppendroff tubes and labelled. It was centrifuged at 5000 rpm for 10 min in a Spinwin™ centrifuge

(Tarsons, India). The supernatant was used for the estimation of total phenolic content.

Estimation of total phenolic content: The total phenolic content in the gall samples were determined by the

Folin-Ciocalteau (FC) method using Gallic acid as a standard (Volluri et al. 2011) with slight modifications.

Different concentration of extracts (50–250 µg ml-1) was mixed with 1.0 ml of Folin-Ciocalteau reagent (1:1

dilution). 2.0 ml of Sodium carbonate (20%, w/v) was added and shaken well. The reaction mixture was

incubated for 45 min under dark. After the specified incubation period the absorbance was read at 765 nm in

UV/Vis Spectrophotometer (T 60, TTL Technologies, India). The absorbance of standard as well as the

samples was plotted. The amount of total phenolics in the samples was expressed in terms of Gallic acid

equivalents (µg ml-1 GAE).

Estimation of total carbohydrate content: The total soluble carbohydrate content in the aqueous extracts of gall

tissues was determined using the phenol sulphuric acid method. Glucose was used as the standard

(Sadashivam & Manickum 2008). The aqueous extracts of the gall was obtained by boiling the known

amount of gall tissue (1 g ml-1). The filtrate was used for the estimation of carbohydrates. A working

solution was prepared by using different concentrations (1 mg ml-1) of standard (5–25 µg). The samples were

aliquoted (50–250 µl). The volumes of aliquots were made 500 µl with distilled water. A blank was set with

500 µl using distilled water. One ml of liquefied phenol solution (5%) was added to all tubes followed by

the addition of 2.0 ml of concentrated sulphuric acid immediately and shaken well. The reaction mixture was

incubated for 30 min in dark place. The absorbance of the standard as well as the samples were read at 490

nm using Spectrophotometer. The absorbance values and the amount of total carbohydrate in the different

samples was calculated and represented.

RESULTS

Identification of the gall on stem/twigs of Michelia champaca L.

Morphological observations: Gall on M. champaca occurred on the main stem axis, as well as twigs, two feet

above the ground (Fig. 1B). The gall during initial development stages (young) appeared as snow-white out

growths (0.6 cm in diameter). They were formed at regular distance both on the stem as well as the twig

(Fig. 1C), as they matured, an increase in size (2.0–2.5 cm in diameter) was observed and later coalesced to

form large sub-spherical structures. One of the important observations made during the gall development is

the change from the initial snow-white colour (Fig. 1C) to slightly blackish structure at maturity (Fig. 1D).

Observation of mature galls by Steriotrinocular microscopy showed the presence of small black glandular

hairs filled with exudates (Figs. 1E & F).

Identification of the gall-maker: The gall-maker on the stem/ twigs of M. champaca was identified as belonging

to the order-Thysanoptera, which include Thrips. Upon periodical and successive observations it was

confirmed as the genus Podothrips. On the basis of the characters, and the availability of various stages of

development (Fig. 2), only one insect chamber was observed from the under surface of the gall. The body

size observed was 0.5–1.0 mm. The systematic position is as follows: Class: Insecta; Order Thysanoptera;

Family: Phlaeothripidae; Genus: Podothrips.

Histochemical studies: A comparison of section of galled, gall normal and healthy portion showed remarkable

differences anatomically (Table 1). In healthy portion, a conspicuous pith and sub-spherical outline of

vascular bundles with the larger bundles alternating with the smaller bundles was observed (Fig. 3A-a). In

gall normal, pith is conspicuous and the outline of vascular bundles is sub-spherical, the larger bundles

alternate with smaller bundles (Fig. 3A-b). The section resembles the healthy plant. In galled portion,

inconspicuous pith, radial growth of vascular bundles and conspicuous sclerenchyma cap was observed (Fig.

3A-c). The following are the results of the tests.

Alkaloids: A golden yellow colour revealed the presence of alkaloids (Figs. 3B-a–c). In healthy tissue,

sclerenchyma cap and metaxylem and vascular bundles and entire pith region were lightly stained. In gall

normal stem tissue, an intense stain was observed in sclerenchyma cap and cells of metaxylem showed less

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intensity of stain and in pith region only centre portion were stained. In gall stem tissue, the sclerenchyma

cap, vascular bundles, medullary rays and metaxylem cells showed the high intensity of stain and outer cells

of the pith region was stained less intensely.

Figure 1. Galls on M. champaca L.: A, Healthy plant; B, Galls observed two feet above the ground level; C, Stem and twigs

showing galls; D, Close up indicating slightly blackish colour; E, Increased size with black glandular hairs filled with

exudates; F, Close up.

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Figure 2. Developmental stages of the gall-maker: A, Healthy plant; B, Infected plant; C, galls on twig; D, Single gall; E,

Dorsal view of gall; F, Ventral view of gall; G, Young thrip; H, Adult thrip.

Fats: Blue colour indicated the presence of fat. In healthy tissue, high intensity stain was observed in

sclerenchyma cap, vascular bundles and medullary rays and less intensity in pith cells, cortex and epidermis.

In gall normal stem tissue, sclerenchyma cap is continuous and forming a ring like structure. In these

vascular bundles, sclerenchyma cap and cortex showed the high intensity and less in pith cells. In galled

stem tissue, sclerenchyma cap was discontinuous. Sclerenchyma cap, medullary rays and vascular bundles,

showed the high intensity of stain, very less in pith regions (Figs. 3C-a–c).

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Tannins: Blue colour indicated the presence of tannins. In healthy tissue, in xylem elements dark stain was

observed, high intensity of stain was observed in the sclerenchyma cap, vascular bundle and medullary rays.

The pith region showed very high intensity of stain. In gall normal stem tissue, the sclerenchyma cap,

vascular bundle and medullary rays showed the high intensity of stain and in pith region showed the very

high stain. In galled tissue, except the sclerenchyma cap all cells showed the high intense stain (Figs. 3D-a–

c).

Table 1. Histochemical differences in the healthy and galled twig of M. champaca L.

Histochemical

tests Color

Histological components stained

Healthy twig portion Galled twig portion

Alkaloids Golden

yellow

Sclerenchyma cap, vascular bundles &

entire pith (+)

Sclerenchyma cap, vascular bundles,

medullary rays (+++) & outer pith (+)

Fats Cobalt

blue

Sclerenchyma cap ( Continuous ring),

vascular bundles, medullary rays

(+++); pith cells, cortex, epidermis (+)

Discontinuous sclerenchymatous cap,

vascular bundles, medullary rays (+++)

Tannins Prussian

blue


medullary rays & pith region (+++);

Sclerenchyma cap (+); vascular bundles,

medullary rays & pith region (+++)

Starch Blue Cortex, medullary rays & pith (+++); Sclerenchyma cap, metaxylem elements

& pith region (+++); cortex (+)

Proteins Intense

yellow


medullary rays & pith region(+)


medullary rays (+++);pith region (+)

Note: +, indicates light staining; +++, indicates moderate staining; +++, indicates intense staining.

Starch: Blue colour indicated the presence of starch, in healthy tissue, high intensity starch was observed in

cortex, medullary rays and pith cells, pith is conspicuous (Fig. 3E-a). In gall normal tissue, cortex,

metaxylem cells, medullary rays showed high intensity of stain. In pith region very high intensity of stain

was observed (Fig. 3E-b). In galled tissue, sclerenchyma cap, metaxylem elements and pith region showed

the intense stain, more in vascular region and less in cortex (Fig. 3E-c).

Protein: An intense yellow colour indicated the presence of proteins; almost all the tissue showed intense

staining. In healthy tissue, less intensity of stain was observed in the sclerenchyma cap and very less in all

parts of the sections (Fig. 3F-a). In gall normal tissue, high intensity of stain was observed in sclerenchyma

cap and metaxylem cells and less in vascular bundles and medullary rays and pith region showed less

intensity (Fig. 3F-b). In galled tissue, sclerenchyma cap, metaxylem elements, medullary rays and vascular

bundles showed the very high intensity and less intensity in the pith region (Fig. 3F-c).

Total phenolic and total soluble carbohydrate contents

Total phenolics: The phenolic contents of young and mature gall tissue having differences in the growth stages

were compared with Gallic acid as standard. In young and mature galls, the phenolic content of 240 µg ml-1

GAE and 225 µg ml-1 GAE were observed (Fig. 4). Both did not differ much in their phenolic contents.

However, a 2.3-fold difference was noted, when expressed in terms of weight of the gall tissue.

Total soluble carbohydrates: The total soluble carbohydrate content of young and mature gall tissues showing

much difference in the growth stages were compared with glucose as standard. The total soluble

carbohydrate was 9.4 µg ml-1 in young galls, whereas the mature gall contained 0.5 µg ml-1 (Fig. 5).

However, an 18.8-fold increase was noted in the concentration in the young galls when compared to mature

galls.

DISCUSSIONS

Insect galls are organized structures of plant tissue which form in response to either feeding by the gall-

maker (phytophagy) or egg-laying (oviposition) on plant tissue. There are number of galls types induced by the

insects on leaf, stem, twig, bud or flower and fruit. In the present investigation, galls on Michelia champaca

(Magnoliaceae) were noticed for the first time on the stem/twigs. So far, a stipular scar forming a ring around

the stem (node) in Magnolia virginiana has been reported to be caused by a psyllid, Trioza magnolia (Ashmead)

(Hall 2009).

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Figure 3. A, Healthy sections stained with safranin stain; Histochemical tests for: B, Alkaloids; C,

Proteins; D, Fats; E, Tannins; F, Starch (a, Healthy portion; b, Gall normal portion; c, Galled portion).

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Figure 4. Determination of total phenolic content in the gall tissues of M. champaca; Ethanolic extracts of young and mature

gall tissues were prepared (1g 10 ml-1). Concentrations ranging from 50-250 μg ml-1 were tested for the total phenolic

content by Folin-Ciocalteu method using gallic acid as standard. The absorbance of the samples as well as the standard was

read at 765 nm and the values of the samples were calculated from the standard graph and represented in terms of μg ml-1

GAE.

Figure 5. Determination of total soluble carbohydrates in gall tissues of M. champaca: The total carbohydrate content of the

aqueous extracts of young and mature galls were determined by phenol-sulphuric acid method using glucose as standard.

Concentrations ranging from 50-250 μg ml-1 were tested for the carbohydrate content. The absorbance of the samples and the

standard was read at 490 nm and the values of the samples were calculated from the standard graph and represented.

Galls are made up of cells that are more numerous or larger than the normal plant cell or plant organs whose

growth and development have been altered into unusual shapes. An insect gall is initiated because of the plant’s

response to the insect’s egg laying (oviposition) or presence of the egg, and/or feeding stimulation by the larva

(phytophagy). Plant cells are usually modified and enlarged, the plant tissue surrounds the egg or larva, and the

gall protects and feeds the gall-inducer. Galls are located on rapidly growing plant parts-on catkins, seeds,

flowers, petioles, branches and stems; most occur on leaves and buds. Some galls are single-chambered

(monothalamous) and contain only one gall-maker, and others are multi-chambered (polythalamous) which

contain many gall-makers (Buss 2011).

Gall on M. champaca occurred on the main stem axis, as well as lateral twigs, two feet above the ground.

The gall during initial development stages (young) appeared as snow white out growths (0.6 cm in diameter).

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They were formed at regular distance and as they matured galls increased in size (2.0–2.5 cm diameter) and

coalesced to form large sub-spherical structure. Gopinathan & Suresh (1985) worked on the solid, indehiscent

galls on the stem of Pongamia glabra induced by an undescribed agromyzid (Diptera). They reported, that

young galls (8–15 × 6–12 mm) were relatively soft and green and with maturation, they became harder and

developed white patches and is directly proportional to the number of larvae residing inside. Mature galls

showed generally, a single gall chamber extending along the pith region.

The gall-inducer was identified as belonging to the genus Podothrips of the order- Thysanoptera, family–

Phlaeothripidae. Only a single insect chamber was observed from the under surface of the gall. The body size

observed was 0.5–1.0 mm. The genus Podothrips are reported for the first time as gall-inducers. Furthermore,

thrips as gall-makers have been reported on plant species (Raman & Ananthakrishnan 1989). Thrips such as

Gynaikothrips uzeli induced galls on the leaves of Ficus bengalensis (Borbon 2011). Gynaikothips ficorum

caused leaf gall on Ficus laevigata (Santis 1980). Although, we have documented the evidence of this genus as

a gall-inducer, worldwide Podothrips are reported to cause huge losses to crops such as chilli (Scirtothrips

dorsalis), which is an important pest of crops in tropical and subtropical regions in Florida (Ludwig & Bogsan

2007).

A comparison of sections of galled, gall normal and healthy portion showed remarkable differences

anatomically. The following tests were conducted to localize viz., alkaloids, fats, starch, tannins and proteins in

tissues. In healthy portion, pith was conspicuous and the outline of vascular bundles was sub-spherical, the

larger bundles alternated with the smaller bundles. In gall normal, pith is conspicuous and the outline of vascular

bundles is sub-spherical, the larger bundles alternate with smaller bundles. The section resembles the healthy

plant. In the galled portion, inconspicuous pith, radial growth of vascular bundles and conspicuous

sclerenchyma cap was observed.

In the present study alkaloids, tannins, fat, proteins and starch were detected in various tissues and differed

considerably in healthy and galled portions of the sections. Our observations are supported by the studies of

Gopinathan & Suresh (1985) who worked on solid, indehiscent galls on the stem of Pongamia glabra induced

by an undescribed agromyzid (Diptera). In the histochemical studies, maximum concentrations of proteins were

observed in the nutritive cells of medullary region and phloem cells. Starch deposits occurred in the pith cells,

medullary rays in normal stem sections, whereas in gall portions they were located in the outer region of pith of

the nutritive zone. Tannins were present less in gall tissues, while they were totally absent in normal stem.

Kumar & Mathur (2009) reported the histochemical localizations in stem gall of Terminalia arjuna caused by

unknown Itonididae (Diptera). According to the authors, in normal stem sections, cortex, medullary rays and

pith cells contained starch as high intensity of staining was observed. It was present in high quantity in nutritive

zone and vascular region and less in cortical parenchyma cells of stem gall tissue. High intensity of proteins was

observed in the cortical zone. In normal stem, tannins were observed in cork, cortex, vascular and pith regions.

High amount of tannins were observed in cortical region and tannin filled cells were observed near the nutritive

zone of the stem gall. High protein was found in gall tissue as compared to normal tissue from the leaf gall of

Pongamia pinnata induced by Aceria pongamiae (Kumar 2012). The presence of starch near the gall cavity

suggests that the insects are utilizing starch as food materials as such. Increased amount of proteins were noticed

in the nutritive tissues, which helps the gall-makers growth and development.

An increased amount of tannins in gall tissue could be attributed to the higher incidence of polyphenol

activity. Phenolics are one of the much interest in phytochemical as bioactive components of food. It is now

possible to establish the antioxidant activities of plant derived phenolics in the aqueous and lipophilic phases.

The inner walls of the galls are moist and are usually rich in phenolics and carbohydrates. In the present study,

phenolic contents of young and mature gall tissue with differences in the growth stages were compared. In

young galls, 240 µg ml-1 GAE phenolic content and 225 µg ml-1 GAE were observed in mature galls, suggesting

fewer differences in the content among young/mature gall tissues. Ramani & Kant (1989) reported that phenolic

content in the galls of Prosopis cineraria (L.) induced by Lobopteromyia were higher in the normal compared to

gall tissue both in vivo and in vitro. Abrahamson et al. (2003) investigated that rapidly growing galls in three

additional susceptible clones confirmed the increase in phenolics and showed that phenolic level increases as

much as five-fold in galls near their peak growth period. Akhlaq & Mohammed (2011) reported that the stem

gall of Tamarix aphylla was rich in phenolic compounds exhibited antioxidant activity.

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The total soluble carbohydrate content was 9.4 µg ml-1 in young galls, whereas the mature gall contained 0.5

µg ml-1, which is negligible. However, 18.8-fold increase was noted in the concentration in young galls when

compared to mature galls. It may be inferred that high levels of carbohydrates are required during early stage of

gall development as a source for the development of insects. Still, more sampling and assessment of antioxidant

activity in the extracts may reveal that galls are needed good sources of antioxidants. To conclude, the gall

serves both as a shelter and a food source, also the gall-maker is partially protected from parasites and predators.

Further investigation to identify the species of the gall-maker of M. champaca would be beneficial, to elucidate

the life cycle as well as ecological studies.

ACKNOWLEDGEMENTS

The authors are grateful to the Chairman, DOS in Botany, and DOS in Zoology, University of Mysore,

Mysore, Karnataka, India for providing the necessary facilities during the study. The paper is the sincere

outcome of the second author KES during the MSc project work at the DOS in Botany, University of Mysore.

This work has been supported by funds from the University of Mysore.

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Documents

Stem gall of Michelia champaca L. (Magnoliaceae) … flower locally known as ‘sampige’ or ‘champa’ has great significance in Hindu mythology and has a number of cosmetic, medicinal